47

CHAPTER 2-5
PEATLAND RHIZOPODS

Figure 1. A peatland with Sphagnum magellanicum that serves as habitat for protozoa. Photo by Michael Lth.

Peatlands Taxa: Sphagnum

Protozoa, and especially Rhizopoda, are apparently
most abundant in peatlands (Figure 1) and were among the
earliest of the moss fauna to be examined (Jung 1936). But
few other bryophyte protozoans have been studied in detail.
Among the abundant sphagnicolous species (growing in
Sphagnum moss) are Nebela (Figure 2), Hyalosphenia
(Figure 3), Difflugia pyriformis (Figure 4), and D.
globularis (Bovee 1979; Gerson 1982).

Figure 3. Hyalosphenia elegans, a sphagnicole. Photo by

Yuuji Tsukii.

Figure 2. Nebela collaris, a sphagnicole. Photo by Yuuji

Tsukii.

Mitchell et al. (2000b) compared testate amoebae in

peatlands of Switzerland, the Netherlands, Great Britain,
Sweden, and Finland. They found that the plant species
differed more than the species of amoebae. The high
number of rhizopod species among Sphagnum, compared
to that of other mosses or tracheophytes, supported the
usefulness of rhizopods as indicators of both past and
present conditions. Furthermore, the mosses were less

48

Chapter 2-5: Peatland Rhizopods

affected by the chemistry of the ground water than were

such taxa as Carex and Eriophorum. But when Booth and
Zygmunt (2005) compared the testate amoeba communities
of the Great Lakes in North America with those of the
Rocky Mountains of North America, the communities
differed, perhaps due to differences in climate and the
trophic state of the peatlands. Even so, these two regions
had many species in common, and these species occupied
similar moisture positions in both regions. In the Rocky
Mountains, USA, distribution of these testate amoebae in
Sphagnum-dominated peatlands is dictated primarily by
surface moisture (Zygmunt et al. 2003). Communities in
the western Great Lakes region are similarly distributed,
with 50% of the species also occurring in the Rocky
Mountain peatlands, and similar communities exist for
Yellowstone National Park.

ecology than by geography. This is reflected in the smallscale vertical gradients seen among the amoebae, rotifers,
and other invertebrates. As noted above, it appears that the
number of species of these rhizopods is generally much
greater among Sphagnum (Figure 1) than among other
mosses or tracheophytes (Mitchell et al. 2000b).
Nevertheless, Tolonen et al. (1992) found little difference
in rhizopod taxa between Sphagnum communities and
those of bryalean mosses in Finnish mires. Unfortunately,
few studies have compared fauna on these two groups of
bryophytes at the same location.

Figure 5. Assulina muscorum showing pseudopodia and test.

Photo by Yuuji Tsukii, Protist Information Server.
Figure 4. Difflugia pyriformis, a sphagnicole. Photo by
Yuuji Tsukii, Protist Information Server.

Testate amoebae abound in peatlands all over the

world. Because of their abundance there, testate amoebae
have been widely studied in peatlands all over the world
(e.g. Leidy 1879; Harnish 1924, 1925, 1927, 1948, 1950,
1951; Hoogenraad 1934, 1935; Jackzo 1941; van Oye
1941, 1951; Conra, 1943; Heinis 1945; Hoogenraad & de
Groot 1946; Paulson 1953; Rose 1953; Hoppman 1954;
Chacharonis 1956; Varga 1956; Bonnet 1958; Thomas
1959; Heal 1961, 1964; Schnborn 1962, 1963, 1965;
Martin 1963; Buttler et al. 1966 a, b; Tolonen 1966, 1994;
Coteau 1969; Bovee 1979; Seis 1971; Corbet, 1973;
Laminger 1975; Vucetich 1975; Grospietsch 1976;
Ruitenburg & Davids 1977; Meisterfeld 1978, 1979a, b;
Beyens & Chardez 1984; Tolonen et al. 1985, 1992, 1994;
Warner 1987; Hendon & Charman 1997; Gilbert et al.
1998a, b, 2003; Woodland et al. 1998; Bobrov et al. 1999;
Strder-Kypke & Schnborn 1999; Mitchell et al. 1999,
2000a, b; Charman et al. 2000; Booth 2002; Langdon et al.
2003; Laggoun-Dfarge et al. 2008).
Bobrov et al. (1999) studied their ecology in peatlands
of Russia. Bousquet (1950) studied them in southwestern
France, Mieczan (2006) in Poland, and Wilmshurst (1998)
in New Zealand. Robson et al. (2001) reported on
Sphagnum bog microfauna in Tierra del Fuego, South
America, demonstrating several of the same familiar genera
as those in Switzerland (Bartos 1949a). Among those
Northern Hemisphere taxa also identified in Tierra del
Fuego were Assulina (Figure 5), Corythion (Figure 6),
Euglypha (Figure 7), and Heleopera (Figure 8). Just as
peatland plants are more cosmopolitan than other plants,
these rhizopod assemblages seem to be more affected by

Figure 6. Corythion pulchellum showing lower surface.

Photo by Yuuji Tsukii, Protist Information Server.

Figure 7. Euglypha test sitting on algal filament. Photo by

Jason Oyadomari.

Chapter 2-5: Peatland Rhizopods

Figure 8. Heleopera sp. test with protoplast. Photo by Yuuji

Tsukii, Protist Information Server.

The nature of peatlands may account for their

prominent testate amoeba fauna (Booth & Zygmunt 2005).
Sphagnum itself is particularly rich in species (Hingley
1993; Mazei et al. 2007). The amoebae are able to live in
the thin film of water in the concavity of Sphagnum leaves
(Figure 9; Corbet 1973). Mazei et al. (2007) found 59
species of testate amoebae among the Sphagnum plants of a
bog in Volga Highland in Russia. Among these, 24 were
common and the minimal richness was three species in a
sample. Interestingly, the highest densities of organisms
occurred in the driest bog habitats, but predictably, the
diversity was lowest (3 species), with Arcella arenaria
(Figure 10) the most common. At medium levels of
humidity, the number of species was greater (13-16), with
Nebela tenella (Figure 11) and Hyalosphenia elegans
(Figure 12) being the most common. Low oxygen
When
concentrations reduced densities by 50-65%.
oxygen was not limiting, however, both abundance and
species richness increased with depth. At high humidity,
the dominant taxa were Hyalosphenia papilio (Figure 13)
and Heleopera sphagni (Figure 14). But not all of these
testae were occupied by live amoebae. The number of
living individuals ranged 35-75% of the testae found.

49

Figure 10. Arcella arenaria test. Photo by Yuuji Tsukii,

Protist Information Server.

Figure 11. Nebela tenella test with protoplast. Photo by

Yuuji Tsukii, Protist Information Server.

Figure 12. Hyalosphenia elegans test. Photo by Edward

Mitchell.
Figure 9. Sphagnum papillosum showing the hood leaf tips
that provide a concavity for water that houses amoeboid protozoa.
Photo by Michael Lth.

Lamentowicz and Mitchell (2005) found 52 taxa of

testate amoebae in Sphagnum peatlands of northwestern
Poland. In a later study, in Poland's largest peatland
complex, Lamentowicz et al. (2007) found 32 taxa of
testate amoebae. In most of the ten sites in this complex,
species composition was dominated by Hyalosphenia
papilio (Figure 13), Cyclopyxis arcelloides (see Figure 15),
and Hyalosphenia elegans (Figure 12); Amphitrema flavum
(Figure 16, Figure 17) was among the most numerous.

50

Chapter 2-5: Peatland Rhizopods

Corbet (1973) found several species that are apparently

confined to the Sphagnum habitat: Amphitrema flavum
(Figure 16, Figure 17), A. wrightianum (Figure 18, Figure
19), A. stenostoma (Figure 20), Hyalosphenia elegans
(Figure 12), and H. papilio (Figure 13). Cryptodifflugia
ovalis (Figure 21) and Amphitrema flavum (Figure 16) can
live within the hyaline cells of Sphagnum leaves.

Figure 13. Hyalosphenia papilio test. Photo by Edward

Mitchell.

Figure 16.
Several species, such as this rhizopod
[Amphitrema (=Archerella) flavum] are confined to the Sphagnum
habitat. It is shown here in a Sphagnum leaf. Photo by Edward
Mitchell, 2004. From Genome News Network, The Wet World of
Moss
<http://www.genomenewsnetwork.org/articles/2004/03/04/moss.p
hp>.

Figure 14. Heleopera sphagni living cell and test. Photo by

Yuuji Tsukii, Protist Information Server.

Lamentowicz and Mitchell (2005) identified three

groups of testate taxa, based on depth to water table (DWT)
and pH: high DWT & low pH, low DWT & low pH, and
high pH & mid-range DWT. It appears that species
tolerance increases with dryness, with a pattern that reflects
that of Sphagnum. That is, changes in the water table depth
have more effect on those species in wet habitats than on
those in drier microhabitats. This appears to indicate that
those in dry microhabitats are specialists for drought..

Figure 17.
Amphitrema (Archerella) flavum showing
pseudopods. Photo by Edward Mitchell.

Figure 18. Amphitrema wrightianum showing ingested

chloroplasts. Photo by Edward Mitchell.

Figure 15. Cyclopyxis, a testate rhizopod. Photo by Yuuji

Tsukii, Protist Information Server.

Chapter 2-5: Peatland Rhizopods

Figure 19. Amphitrema wrightianum using fluorescence to

show ingested chloroplasts. Photo by Edward Mitchell.

Figure 20. Amphitrema stenostoma test with sand grains and

living protoplast with included chloroplasts. Photo by Yuuji
Tsukii, Protist Information Server.

Figure 21. Cryptodifflugia ovalis showing living cell and

extruded protoplasm. Photo by William Bourland.

Those species that characterized Sphagnum hummocks

(Figure 22) in the western Carpathians [Nebela militaris
(Figure 23), N. tincta (Figure 24), Assulina muscorum
(Figure 25), Heleopera petricola (Figure 26)] seemed
intolerant of the mineral-rich fens (Opravilov & Hjek
2006). Only Corythion dubium (Figure 27) and Nebela
bohemica seemed able to occupy both. The Euglyphidae
were dominant in all these habitats and were nearly the
exclusive testate inhabitants of the moderately rich fens.
Hyalospheniidae, on the other hand, characterized the
extremely acid habitats, particularly in Sphagnum
hummocks. The overall vegetation was the best predictor
of the testate protozoan composition, and the composition
of the bryophyte assemblage was the second most
important predictor.

51

Figure 22.
Michael Lth.

Sphagnum warnstorfii hummock.

Figure 23.
Mitchell.

Test of Nebela militaris.

Photo by

Photo by Edward

Figure 24. Nebela tincta test. Photo by Yuuji Tsukii, Protist

Information Server.

Figure 25. Assulina muscorum test. Photo by Yuuji Tsukii.

52

Chapter 2-5: Peatland Rhizopods

Submerged Sphagnum riparium is characterized by an

association of Cyclopyxis eurystoma, Heleopera sphagni,
Hyalosphenia papilio, and Phryganella hemisphaerica.
Available moisture, determined by depth from the water
table, separated the communities.
The greatest
homogeneity occurs in the moist areas in the middle of the
quagmire, whereas the dry habitats have the greatest
diversity. On the other hand, a greater proportion of the
amoebae were alive in the moist areas (36-45%) compared
to 22-27% of those in dry habitats.

Medium and Rich Fens

Figure 26. Heleopera petricola test beside a desmid. Photo

by Yuuji Tsukii.

Bryophytes of rich fens (Figure 28) differ greatly from

those of Sphagnum bogs and poor fens, and so do the
protozoa.
To utilize fully the testate protozoa to
reconstruct peatland history, as discussed later in this
chapter, it is important to understand these faunal
differences. Opravilov and Hjek (2006) studied the
spring fens of the Western Carpathians in the Czech
Republic and Slovakia to fill in this rather large gap in our
knowledge. They found that two species [Paraquadrula
irregularis (Figure 29, Figure 30) and Centropyxis
discoides (see Figure 31)] were essentially restricted to
fens, while seven rhizopod species characterized the
bryophytes there. In moderately rich Sphagnum fens,
Arcella discoides (Figure 32) was characteristic. In poor
fens, testate protozoan species of bryophyte lawns were
closely tied to moisture and overlapped widely with those
of poor fen sediments and moderately rich fens: Nebela
collaris (Figure 33), Phryganella acropodia, Sphenoderia
fissirostris.

Figure 27. Corythion dubium. Photo by Yuuji Tsukii.

Mazei and Tsyganov (2007/08) reported on a number

of taxa in the Sphagnum peatlands of Russia. In a single
bog, they found 63 taxa comprising 21 genera. They found
two different communities, one that lived in the Sphagnum
"quagmire" and one that lived in the bottom sediments of
the drainage. The detritivores from the bottom sediments
included Arcella gibbosa, A. vulgaris, A. hemisphaerica, A.
discoides, A. intermedia, A. mitrata, Centropyxis aculeata
sphagnicola, Cyclopyxis kahli, Difflugia glans, Lesquereusia
spiralis, Netzelia tuberculata, and Phryganella hemisphaerica.
Those species typical of Sphagnum were Archerella flavum,
Euglypha cristata, Difflugia juzephiniensis, Cryptodifflugia
compressa, Nebela militaris, and Sphenoderia fissirostris.
Those inhabiting both the Sphagnum mats and the quagmire
inclued Assulina seminulum, A. muscorum, Bullinularia
indica, Centropyxis aculeata, Difflugia globulosa, D. parva,
Euglypha ciliata, Hyalosphenia elegans, Nebela tenella, and
N. tincta. Other species are not so specific and occur in both
of the major bog communities: Arcella arenaria, Euglypha
laevis, and Trigonopyxis arcula.
But even within the Sphagnum quagmire, Mazei and

Tsyganov (2007/08) found three types of testate amoebae

communities. The xerophilous community could be found
in hummocks made of Polytrichum strictum, Sphagnum
papillosum, and S. angustifolium. These dry hummocks
house a community characterized by Assulina muscorum,
A. seminulum, and Cryptodifflugia compressa. The lawns
of Sphagnum palustre and S. magellanicum make a wet
community characterized by Heleopera sphagni,
Hyalosphenia papilio, H. elegans, and Nebela tenella.

Figure 28. Limprichtia (=Drepanocladus) revolvens in a rich

fen. Photo by Michael Lth.

Figure 29. Paraquadrula sp. showing test.

Edward Mitchell.

Photos by

Chapter 2-5: Peatland Rhizopods

Figure 30. Paraquadrula irregularis. Photo by William

Bourland.

Figure 31. Centropyxis ecornis. Photo by Yuuji Tsukii,

Protist Information Server.

Figure 32. Arcella discoides test and protoplast. Photo by

Yuuji Tsukii, Protist Information Server.

53

Figure 33. Nebela collaris test and cell. Photo by Yuuji

Tsukii, Protist Information Server.

The protozoan species of Sphagnum fens in the Czech

Republic and Slovakia are very similar to those known
elsewhere, with Amphitrema flavum (Figure 34), A.
wrightianum (Figure 34), and Hyalosphenia papilio (Figure
35), being optimal in wet microhabitats, but also tolerating
higher mineral concentrations (Meisterfeld 1979b;
Charman & Warner 1992; Tolonen et al. 1992; Booth
2001; Schnitchen et al. 2003; Booth & Zygmunt 2005;
Lamentowicz & Mitchell 2005; Opravilov & Hjek 2006).
In the drier poor fens, the dominant species are Assulina
muscorum (Figure 25), A. seminulum (Figure 36), Arcella
catinus (Figure 37), Nebela militaris (Figure 23), N.
bohemica, Trigonopyxis arcula (Figure 38), and Corythion
dubium (Figure 39). Corythion dubium also occurs in
moderately rich fens (Beyens et al. 1986; Tolonen et al.
1994; Bobrov et al. 1999; Mitchell et al. 2000b; Opravilov
& Zahrdkov 2003; Vincke et al. 2004).

Figure 34. Left: Amphitrema flavum. Right: Amphitrema

wrightianum. Photos by Edward Mitchell.

54

Chapter 2-5: Peatland Rhizopods

Figure 35. Hyalosphenia papilio. Photo by Yuuji Tsukii.

Figure 39. Test of Corythion dubium.
Tsukii, Protist Information Server.

Figure 36. Assulina seminulum. Photo by Yuuji Tsukii.

Photo by Yuuji

Among the "brown mosses" (Figure 40, Figure 41, ) of

calcareous fens, Centropyxis cassis, Cyclopyxis kahli,
Cyphoderia ampulla (Figure 42), Difflugia glans,
Quadrulella symmetrica (Figure 43), and Trinema enchelys
(Figure 44) often predominate (Mattheeussen et al. 2005;
Opravilov & Hjek 2006). There is indeed a gradient of
species from poor to rich fens, with moisture being an
important variable in the poor fens and bogs (Opravilov &
Hjek 2006). Interestingly, the sediments of poor acidic
fens support a species composition similar to that of
bryophyte tufts of mineral rich fens.

Figure 40. Tomentypnum nitens, a brown moss common in

fens. Photo by Michael Lth.
Figure 37. Arcella catinus test. Photo by Yuuji Tsukii,
Protist Information Server.

Figure 38. Trigonopyxis arcula. Photo by Yuuji Tsukii,

Protist Information Server.

Figure 41. Scorpidium scorpioides, a brown moss common

in fens. Photo by Michael Lth.

Chapter 2-5: Peatland Rhizopods

55

drainage that resulted in an increase in tree cover, especially at the

edges. By the second sampling date in 2001, three species had
increased significantly in mean relative abundance: Nebela tincta
s. l. (+97%), Bullinularia indica (+810%), and Cyclopyxis
eurystoma (+100%; absent in 1961), while two species decreased
significantly: Assulina muscorum (-63%) and Euglypha
compressa (-93%).
Furthermore, testate amoebae

Figure 42.
Mitchell.

Cyphoderia ampulla test.

Photo by Edward

communities differed among hummocks, lawns, and

hollows. Nevertheless, there were no significant changes
in the overall community structure between the two
sampling dates.
Recent, moist stages of succession in the Jura
Mountains of Switzerland were dominated by
Hyalosphenia papilio, with Archerella flavum indicating
wet, acidic conditions at one site (Laggoun-Dfarge et al.
2008). Drier acid conditions supported a greater abundance
of Nebela tincta and Assulina muscorum. Corythion
dubium also indicated dry, acid conditions.

Habitat Needs

Figure 43. Quadrulella symmetrica. Photo by Yuuji Tsukii.

Mieczan (2007) examined the habitat preferences of

eleven testate amoebae in Eastern Poland peatlands. He
found that low pH (4.5) favored the amoebae (see also
Warner & Chmielewski 1992; Tolonen et al. 1994;
Charman & Warner 1997; Mitchell et al. 1999; Bobrov et
al. 2002; Booth 2002; Lamentowicz & Mitchell 2005).
These acidophilic taxa were dominated by ubiquitous and
common taxa, with Arcella vulgaris, Assulina muscorum,
Euglypha sp., and Hyalosphenia sp. having a distinct
preference for low pH. The distribution pattern seemed to
be controlled by moisture (no surprise there), whereas the
total numbers and biomass had a positive correlation with
pH and total organic carbon content of the water. Heal
(1964) found that pH was a major factor accounting for
differences between bog and fen communities in Great
Britain. In addition to moisture and pH, the trophic status
and concentration of mineral nutrients, including calcium,
can play a role in determining numbers (Tolonen et al
1992).

Food

Figure 44. Trinema enchelys. Photo by Yuuji Tsukii, Protist

Information Server.

Successional Stages
Differences occur not only between peatlands, but also
in different stages of the same peatland, an important factor
in permitting us to reconstruct the past history of peatlands.
Mazei and Bubnova (2007) demonstrated 42 species in the
initial stage of a transitional bog. Early stages were
characterized by widespread species such as Assulina
muscorum, Arcella arenaria, Phryganella hemisphaerica,
and Euglypha laevis, whereas the sphagnobionts such as
Nebela, Hyalosphenia, and Heleopera were absent.
Vertical differences had not developed because the species
that characterize the different depths had not yet become
established.
Kishaba and Mitchell (2005) carried out a 40-year
study on the Sphagnum-inhabiting rhizopods to determine
successional trends in the Swiss Jura Mountains. They took

their first samples in 1961 following peat cutting and lateral

Although many of the protozoa associated with

bryophytes are detritus/bacterial feeders, some common
species prefer a different diet. In one Sphagnum peatland
17.4% of Nebela collaris sensu lato most frequently preyed
upon micro-algae (45%, with diatoms comprising 33% of
total prey), spores and fungal mycelia (36%), and large
ciliates, rotifers, and small testate amoebae in smaller
numbers (Gilbert et al. 2003). However, 71% of the food
content could not be identified because it was partially
decomposed.
It appears that when the mosses are
sufficiently wet, most of the food organisms are immboile,
senescent, or dead. However, as the water film on the moss
becomes thin, it constrains the ciliates and micro-Metazoa,
causing them to be a more easily consumed part of the diet.

Vertical Distribution
Peatlands have both horizontal and vertical differences
in moisture, light availability, nutrient availability, and pH
(Figure 45). The testate rhizopods are distributed both
vertically and horizontally with respect to these differences
(Meisterfeld 1977).

56

Chapter 2-5: Peatland Rhizopods

Figure 45. Sphagnum teres, demonstrating the zonation from

light to dark within the peat. Photo by Michael Lth.

Perhaps because of the multiple factors involved in

vertical and horizontal distribution, distinct patterns are
difficult to discern.
Mazei and Tsyganov (2007/8)
considered the aggregations of species to blend into each
other in patches of varying sizes. For Assulina muscorum
and A. seminulum, patch size seemed to correlate with shell
size. As sample size increases, heterogeneity increases.
Communities can be distinct on as small as a 1-cm patch, but
more typically the minimum size does not exceed several cm.
In their study in the Middle Volga region of Russia, Mazei and
Tsyganov found that associated with the upper parts of
Sphagnum the typical species were Assulina flavum, A.
muscorum, A. seminulum, Heleopera sphagni, and
Hyalosphenia papilio.
Among these, Assulina flavum,
Heleopera sphagni, and Hyalosphenia papilio were
mixotrophs, requiring light for their algal symbionts (see subchapter 2-4), whereas Hyalosphenia elegans lacked symbionts
and lived in a deeper community. The upper 0-3 cm layer
typically had low rhizopod species richness but the highest
abundance in the peatlands. And among those tests the
proportion of living organisms was highest (75%). Species of
Amphitrema likewise occur in the upper layer because of the
need for light by their symbionts (Gilbert & Mitchell 006).
When conditions are somewhat drier, the vertical
structure of the communities is more pronounced (Mazei &
Tsyganov 2007/08). Low moisture typically resulted in empty
tests, especially in Assulina species. Survival of the rhizopod
species is facilitated by the r-strategies of reproduction in
which these small organisms are able to increase rapidly in
response to the return of favorable conditions.

One additional factor that may play a role in

distribution for some species is available nitrogen (Mitchell
& Gilbert 2004). In cutover peatlands fertilized with N for
three years, richness of the peatland was high (22 taxa of
testate amoebae), but diversity of individual samples was
low (6.6), attesting to the diversity of the habitat. Species
richness increased with depth, but there was little response
to differences in N levels in the tested range of additions of
0, 1, 3, or 10g N m2 yr1 for three years. Only Bullinularia
indica was significantly more abundant in N-fertilized
plots. Although the vertical distributions differed among
species, there seemed to be no relationship to either shell
type or metabolism type. In the top segment (01 cm),
Assulina muscorum was most abundant. At 35 cm
Heleopera rosea, Nebela militaris, and Phryganella
acropodia were most abundant.

It is not surprising that the taxa with zoochlorellae

occur in the green portions of Sphagnum. In Obersee near
Lunz, Austria, the dominant taxa hosting zoochlorellae are
Amphitrema flavum, Heleopera sphagni, Hyalosphenia
papilio (Laminger 1975). Centropyxis aculeata likewise
lives there, but without zoochlorellae. Activity among the
rhizopods extended down to 18 cm, with some of the less
mobile testate species extending to a depth of 45 cm. Some
of the species that lived down to depths of 12 cm were
species that also inhabited forest mosses (Euglypha laevis,
Trinema enchelys, and T. lineare). At 18 cm, several
sediment species of Difflugia occurred (D. amphora, D.
corona, D. acuminata, D. lebes).
Furthermore, the
populations of Centropyxis aculeata exhibited charactistics
of sediment-inhabiting taxa, i.e. tests covered with mineral
particles and no spines.

Horizontal Differences
Not only do the testate amoebae have a vertical
zonation in peatlands, but their horizontal distribution
varies as well, reflecting habitat patchiness (Meisterfeld
1977; Mitchell et al. 2000a; Mazei and Tsyganov 2007/8).
In the Swiss Jura Mountains, spatial structure accounted for
36% of the observed variation. Imbedded in the horizontal
variability, Mitchell et al. found that microtopography
played an important role, indicating that in just 0.25 m2
conditions are not uniform and present a different picture
from that seen on a macroscale. In this case, the horizontal
scale responds to differences in distance from the water
table, whereas vertically within a Sphagnum mat, light,
moisture, and detrital accumulation all differ.
The
horizontal scale also differs in pH and ion concentrations,
both of which are lower on hummocks than in hollows.
These differences in turn cause differences in the bacteria,
fungi, algae, and other protozoa available for food. And
hummock Sphagnum species are usually different from
hollow species, having different morphologies that provide
different sorts of spaces and different abilities to retain
water and detritus.

Seasonal Differences
Communities of protozoa can differ among seasons,
just as moisture and other conditions change in their
habitat. As a result, species richness will fluctuate, as will
abundance. In a Sphagnum bog in the Middle Volga region
of Russia, species richness increases as the vegetation
increases during May to September (Mazei & Tsygano
2007/08). At the same time, evenness and species diversity
have little variation. Species abundance changes are less
well defined seasonally, most likely being more responsive
to available moisture that is not directly tied to season.
Spring brings melting snow in most peatlands (Figure
46), with dormant protozoa awakening as the environment
In spring, dominant
becomes more hospitable.
hygrophilous (water-loving) species in the Middle Volga
region included Heleopera sphagni, Hyalosphenia papilio,
and Nebela tincta (Mazei Tsyganov 2007/08). This
dominance is replaced in summer and autumn by
Hyalosphenia elegans and Nebela tenella.
The
xerophilous (dry-loving) community is slightly different
and the diversity is somewhat greater. In spring, Assulina
muscorum, Heleopera sphagni, and Nebela tincta
dominate, being replaced in summer by a community of

Chapter 2-5: Peatland Rhizopods

Assulina seminulum, Euglypha ciliata, Hyalosphenia

elegans, and Nebela tenella. Yet another community
appears in autumn, dominated by Assulina seminulum,
Cryptodifflugia compressa, and Trigonopyxis arcula.

Figure 46. As the snow recedes, the Sphagnum habitat will

witness the awakening of water-loving protozoa that have
remained dormant throughout the winter. Photo by Andres Filipe
Baron Lopez in Alaska.

Heal (1964) found slightly different species in his

study of six fen and bog sites in Great Britain, but the
patterns were similar. Three species Amphitrema flavum,
Hyalosphenia papilio, and Nebela tincta sensu lato had
peak numbers from May until October. They then either
encysted or died. For Hyalosphenia papilio, light is a
controlling factor because this protozoan typically contains
photosynthetic zoochlorellae (Figure 47). Although many
of these rhizopods can reproduce every eight days by cell
division, field evidence suggests that they have fewer than
ten generations per year. This low number of generations
limits their ability to respond to improved environmental
conditions. These three species thus accounted for a
biomass of 1.0 g m-2 and 30.2 x 106 individuals m-2 in Great
Britain. Nevertheless, Heal found 98 species and varieties
in these six sites with a distribution similar to that found in
northern fens and bogs.

Figure 47. This protozoan, possibly Bryometopus, contains

zoochlorellae. Photo by Yuuji Tsukii.

One mechanism that maintains closely related species

in different niches is their seasonal requirements. For
example, Hyalosphenia papilio is dominant in spring, H.
elegans in summer-autumn. Nebela tincta occurs in spring,
N. tenella in summer. Assulina muscorum appears in
spring, A. seminulum in summer.

57

Pollution
Pollution can alter the peatland rhizopod communities.
Mitchell et al. (2003) found that CO2 enrichment caused a
change in structure, but not in total biomass. Heterotrophic
bacterial biomass increased by 48%, whereas that of the
testate amoebae decreased by 13%. They suggested that
the increase in CO2 may have caused an increase in
Sphagnum exudates that in turn stimulated an increase in
bacterial biomass.
Protozoan communities are also sensitive to other
pollutants (Nguyen-Viet et al. 2008). Testate amoebae on
Sphagnum fallax declined in species richness, total density,
and total biomass with added lead (Nguyen-Viet et al.
2007, 2008). NO2 also caused a decline in diversity, but
not in density in the more heavily polluted city center of
Besanon, France (34.8 9.5 g m-3) compared to the
peripheral area (14.6 4.7 g m-3) (Nguyen-Viet et al.
2004). Paraquadrula irregularis differed dramatically,
being present in all peripheral samples and completely
absent in the city; no other species differed significantly
between the two areas.

Reconstruction of Past Climate

Diatoms and siliceous protozoan plates and scales are
common in peat preparations (Douglas & Smol 2001).
However, these are seldom used in peatland reconstruction
because it is nearly impossible to identify the species from
these fossils. Fortunately, rhizopod tests are often present
in the same samples and require the same preservation
techniques as the diatoms and scales. Since the species are
generally identified by their shells, there has been
considerable recent interest in using these testate shells for
determining the past history of the peatlands.
Both the mosses and the amoebae are well conserved
over time, Sphagnum because of its resistance to decay, and
for testate amoebae it is the unique test (housing) that
likewise resists decay (Meisterfeld & Heisterbaum 1986;
Coteaux 1992). Both can be identified thousands of years
later.
Even fossil evidence supports the richness of the
Sphagnum fauna (Douglas & Smol 1988). Fortunately, the
species are cosmopolitan (Smith & Wilkinson 2007) and
community structure varies little with geography (Mitchell
et al. 2000b; Booth & Zygmunt 2005), differing much less
between geographic areas than does the tracheophyte
community (Mitchell et al. 2000b). Even if species have
diverged into sister species and become endemic (Mitchell
& Meisterfeld 2005), it will often be possible to use these
species complexes as indicators. On the other hand, we
may be plagued by species that have diverged
physiologically without changing morphologically, thus
permitting them to live under different conditions but
without being recognizable as different taxa.
As already implied, the testate amoebae have a
distribution pattern that mimics that of Sphagnum
(Lamentowicz & Mitchell 2005). Wet habitat species of
both are more sensitive to changes in the water table depth
than are those of dry habitats such as hummocks. Species
of dry habitats are more tolerant of desiccation.
Consequently, the testate amoeba shells from the past
permit us to reconstruct the past history of peatlands (van
Geel 1976; Beyens & Chardez 1987; Warner 1991;
Wilmshurst 1998; Bobrov et al. 1999; Charman et al. 1999;

58

Chapter 2-5: Peatland Rhizopods

McGlone & Wilmshurst 1999a, b; Foissner 1999; Mauquoy

& Barber 2002; Schnitchen et al. 2003; Zygmunt et al.
2003; Booth et al. 2004; Gilbert & Mitchell 2006; Payne et
al. 2006; Payne & Mitchell 2007; Mitchell et al. 2008).
Payne et al. (2008) demonstrated that even such diverse
regions as Turkey, North America, and Europe have similar
testate communities. Because of the unique assemblages of
testate amoebae associated with moisture conditions of the
peat mosses worldwide and the effects of climate change
on them, the testate amoebae are useful for reconstructing
past climate.
Surface moisture of bogs (with only precipitation as a
water source), in particular, is controlled by climate.
Reconstruction of the testate amoeba history permits
reconstruction of the historic surface moisture, and that
permits reconstruction of past rainfall. The amoebae are so
fine tuned to the water table that they can help a researcher
to predict the water table within less than 2 cm (Payne &
Mitchell 2007). For example, Hughes et al. (2006) used
testate amoebae to identify fourteen distinct phases of nearsurface water tables in a coastal plateau bog in eastern
Newfoundland, with corresponding time periods beginning
8270, 7500, 6800, 5700, 5200, 4900, 4400, 4000, 3100,
2500, 2050, 1700, 600, and 200 calibrated years BP. The
final drainage of glacial Lake Agassiz accounts for the first
major phase of pool development at 8400 calibrated
years BP, followed by the Ungava lakes ca 75006900 calibrated years BP. From 7500 BP to the present the
reconstructed bog surface water and the stacked ice rafted
debris of the North Atlantic Ocean correlate well. At the
same time, long-term changes in air masses may have been
a contributing factor. Records of "cosmogenic isotope
flux," when compared to the bog surface wetness
reconstruction, suggest that reduced solar radiation presents
a consistent link with increased bog surface wetness during
the Holocene.
But the models are not always so accurate. Payne et
al. 2006) were only able to estimate within 9.7 cm of water
table depth, and that was after exclusion of selected data.
They attributed the less than ideal fit to inaccuracies in
water-table measurements, very large environmental
gradients, and recent climatic change in the study area.
Their pH estimates were only off by 0.2, which is within
the error range of many pH measuring techniques.
Using weighted averaging to model species abundance
as measures of water table depth and soil moisture, Bobrov
et al. (1999) calculated optima and tolerance of species
niches. They found that each group of taxa tends to have a
gradient of hydrological preference. For example, a wet to
dry gradient is exhibited among species of the Trigonopyxis
arcula group: T. arcula var. major > T. arcula > T.
minuta. Likewise, the Assulina-alkanovia group exhibits
wet to dry as A. seminulum >
A. muscorum >
Hyalosphenia elegans and the Trinema lineare group
appears as T. lineare var. truncatum/T. lineare > T. lineare
var. terricola. Interestingly, these species gradients also
follow a large to small size gradient, indicating that small
taxa survive better than large ones under dry conditions. It
appears that having spines is a disadvantage in dry habitats.
Within the genera Euglypha and Placocista the spined
forms (Figure 48) are typical of wetter habitats than are
those with shorter spines or no spines. These relationships
suggest that the most effective use of these rhizopods for

reconstruction of the past water regime is to use the lowest

possible level of identification, i.e. species and varieties.

Figure 48. Placocista spinosa, a rhizopod typical of wet

habitats. Photo by Yuuji Tsukii.

One interesting question that arises is whether these

spined taxa are really different species and varieties, i.e.,
genetically different, or if they represent ecotypes
morphological representations of the microenvironment
where they occur. For example, Laminger (1975) found
that Centropyxis aculeata at greater depths lacked spines
and their tests were covered with mineral particles. To test
the possibility of ecological morphs, Booth (2001)
examined four of the most common taxa in two Lake
Superior coastal wetlands: Arcella spp., Assulina spp.,
Centropyxis cassis type, and the Nebela tincta-parvulacollaris group. Using 74 microsites, they compared testate
amoeba assemblages based on percent moisture, depth to
water table, pH, porosity, depth of living moss, and
associated moss and tracheophyte species. They used such
parameters as test length and aperture diameter for
amoebae from at least 10 microsites. In general, there was
little correlation between morphological variation and
microenvironmental parameters. However, in the Nebela
tincta-parvula-collaris group, the test size correlated
significantly with pH (r2 = 0.68). Booth concluded that
these testate rhizopods are sensitive indicators of waterlevel and pH changes.
Many more studies on testate amoeba ecology have
been conducted in the Northern Hemisphere than elsewhere
(Mitchell & Meisterfeld 2005), making their comparisons
somewhat easier. In the East Carpathian peatlands of
eastern Europe, species such as Amphitrema flavum (Figure
17) and Hyalosphenia papilio (Figure 12) indicate wet
conditions were present (Schnitchen et al. 2003). Assulina
muscorum (Figure 49), Difflugia pulex, and Nebela
militaris (Figure 23) indicate that conditions were dry.
In Sphagnum peatlands of the Rocky Mountains, USA,
surface moisture determines the distribution of fossil
rhizopods (Zygmunt et al. 2003). As suggested by the
ecological studies of Lamentowicz and Mitchell (2005) and
others (Booth & Zygmunt 2005), Booth and Jackson (2001)
could track the history of an ombrotrophic peatland in
northeastern Lower Michigan, USA, through 2800 years of
changes using the moisture preferences of these organisms.
Such fossils as these testae permit us to determine past
changes in water table depth (Warner 1991; Woodland
1998; Woodland et al. 1998). Booth and Zygmunt (2005)

Chapter 2-5: Peatland Rhizopods

further argued that the widespread geographic nature of the

rhizopod relationships makes interpretation of their
community structure widely applicable.
Charman and Warner (1997) used 60 samples from 14
peatlands in Newfoundland, Canada, and found 40 species
that occurred in more than six samples. They used these to
model the relationships between the species and the water
table depth. Species with narrow tolerances provided the
best indicators. These include Amphitrema stenostoma,
Arcella discoides, Cryptodifflugia sacculus, Difflugia
bacillifera, Nebela carinata, Nebela griseola, Nebela
marginata, Quadrulella symmetrica, and Sphenoderia
lenta. Charman and Warner recommend that for most
accurate results modern constructs from wide regions
should be used to interpret the data from peatland cores that
represent palaeoecological time series.
Fortunately, most of the testate amoeba taxa are
cosmopolitan, permitting the studies from the Northern
Hemisphere to be used in less-studied areas such as New
Zealand (Charman 1997; Wilmshurst 1998). In fact,
Charman (1997) modelled the hydrologic relationships of
protozoa and Sphagnum in peatlands of New Zealand and
suggested that "palaeohydrology could be accurately
inferred from fossil faunas."
Schoning et al. (2005) used peatland amoebae to
reconstruct 125 years of peatland amoebae in Sweden.
Unlike the cases in other areas in Europe, the changes in
water table correlated primarily with changes in mean
annual temperature, whereas in most other studies,
precipitation was also an important factor. They caution
that spatial differences must be considered in these historic
interpretations and thus more study is needed on these
influences.

Figure 49. Test of Assulina muscorum. Photo by Edward

Mitchell.

In a Michigan, USA, study, Booth (2002) found that

most of the eleven peatlands he studied had similar testate
assemblages. As in most other studies, depth to water table
was the best predictor of the protozoan assemblages.
Nevertheless, within a given peatland, community
variability
was
correlated
with
environmental
heterogeneity, adding support to the suggestion of
Schoning et al. (2005) regarding spatial considerations.
But the testate amoebae in bog/fen habitats also had distinct
differences in species between May and late summer-early
autumn. Testate amoebae in the swamp community, on the
other hand, had no clear difference in community structure

59

between dates. They attributed these differences to the

more constant water table and moisture conditions in the
swamp.
Warner et al. (2007) add further support to the
importance of considering seasons, particularly for living
rhizopods. In southern Ontario, Canada, the usual factors
of soil water content and water table influenced the
distribution of amoeboid species and these differ with
seasons.
The historical record will not take us back forever. In
their study on bogs in Ontario and Minnesota, Warner and
Charman (1994) found that cores spanning the entire
Holocene era only exhibited rhizopods present in the last
6500 years. They indicated that the fauna changed from
the early rich fens with sedges and brown mosses. At those
early stages, the protozoan communities were dominated
by Cyclopyxis and Centropyxis. By 5000 BP, the habitat
had become Sphagnum-dominated and the predominant
protozoan taxa had shifted to Amphitrema flavum, Assulina
muscorum, Heleopera sphagni, and Hyalosphenia subflava.
As the habitat became drier, taxa again shifted to Nebela
griseola, N. militaris, and Trigonopyxis arcula.
Geographic Differences
Despite a considerable number of studies indicating
usefulness of these organisms, use of testate amoebae to
determine past habitats can at times be misleading.
Harnish examined mires in Central Europe (1927 in
Paulson 1952-53) and in Lapland, North Sweden (1938 in
Paulson 1952-53), and found that the communities were not
similar. Rather, associations from Central Europe did not
exist in raised bogs in Lapland. In fact, the Amphitrema
association existed in Lapland, but in different habitats, not
raised bogs, whereas in Central Europe it was confined to
raised bogs. The Hyalosphenia type was also absent in the
Lapland raised bogs.
Problems in Using Rhizopods
There are caveats in using fossilized amoeba tests to
assess past communities of testate rhizopods. Not all tests
are equally preserved (Mitchell et al. 2007).
The
Euglyphida, which includes the common Euglypha species
(Figure 50), are an idiosome group that secretes its own
test and its biosilica plates (Beyens & Meisterfeld 2001).
This biological test decays more readily than the testae of
the other groups (Mitchell et al. 2007). In Sphagnum
peatlands, this differential decay seems to make little
difference in the estimations of water table depth.
However, in minerotrophic peatlands, with large numbers
of this Euglyphida group, the loss of these tests leads to an
underestimation of the water table depth. Data on more
alkaline fens are lacking, and the community structure there
is not well known. If this idiosome group is not dominant
there, reconstruction may be more accurate.
Swindles and Roe (2007) likewise found that under
conditions of low pH, such as found in peatlands, the
degree of dissolution was highly variable, but it did not
seem to relate to xenosomic (using "foreign" materials) vs.
idiosomic tests. Euglypha (Figure 50) is particularly
susceptible, whereas Assulina muscorum (Figure 49),
Amphitrema flavum (Figure 34), and Trigonopyxis arcula
(Figure 51) are affected little by acidity. Payne (2007)
found similar results by subjecting rhizopod tests to weak
acid, nutrient enrichment, and desiccation over 28-months,

60

Chapter 2-5: Peatland Rhizopods

and used shorter-term experiments with stronger acids in

peatlands. He determined that during dry periods the
record may be altered by differential preservations of the
tests, as demonstrated by significant effects of long-term
desiccation and short-term acid treatment at two different
concentrations. This consequence could lead to overestimating water table depths.

Figure 50.
SEM detail of biosilica plates of Euglypha
penardi, a protozoan for which the test is especially susceptible to
dissolution. Photo by Edward Mitchell.

In a Polish peatland, a rapid shift in peat accumulation

and lower pH occurred ~110-150 years ago, with a shift to
a Sphagnum-dominated poor fen (Lamentowicz et al.
2007). The protozoa supported this history. Researchers
interpreted this to be a result of forest clearance in
surrounding areas. Whereas peatlands are often destroyed
by human activity, in some cases those activities make
conditions more favorable to peatland development. In this
case, Sphagnum peatland replaced a species-rich poor fen.
Laggoun-Dfarge et al. (2008) found testate amoebae
can be used to reflect disturbances that result from peat
harvesting. Where better carbohydrate preservation was
present, along with more heterogeneous peat composition,
the testate amoebae exhibited a higher diversity, thus
serving as a biological indicator of conditions.
Use in Peatland Regeneration
Regeneration of peatlands sometimes makes use of
remains of testate amoebae to determine the species to reintroduce or to follow the progress in a less labor-intensive
fashion by monitoring the amoebae. In the Jura Mountains,
Switzerland, Laggoun-Dfarge et al. (2008) examined a
peatland that had been mined for heating fuel until World
War II and found that the amoeba communities changed as
the peatlands changed during regeneration. The Sphagnum
habitat shifted from moderately acidic, wet conditions to
more acidic, drier conditions. During these changes, the
biomass and mean size of amoebae declined while
remaining higher at the undamaged site. At the same time,
their species richness and diversity increased while density
declined. As reported by Mitchell et al. (2004), the
changes in the amoeba community lagged behind that of
the returning Sphagnum community. Moreover, during the
forty years of 1961-2001, the overall amoeba richness (33)
remained unchanged, but the richness per sample decreased
from 11.9 to 9.6 (Kishaba & Mitchell 2005). The relative
abundance changed, with three species increasing
significantly [Bullinularia indica (Figure 52) (+810%),
Cyclopyxis eurystoma (+100%, 0 in 1961), Nebela tincta
(Figure 53) (+97%)] and two species declining [Assulina
muscorum (Figure 49) (-63%), Euglypha compressa
(Figure 54) (-93%)]. The researchers concluded that the
expected changes in richness were completed before the
1961-2001 study began.

Figure 51. Trigonopyxis arcula test showing opening for

pseudopod. This test is more stable than that of Euglypha. Photo
by Yuuji Tsukii.

Human Influence on Development

In New Zealand, it appears development of Sphagnum
bogs has been dependent on human activity such as
clearing or modifying the vegetation, resulting in
Sphagnum dominance (Wilmshurst 1998). In other places,
clearing of a peatland means that without human
intervention it is gone forever. After such loss, it is often
desirable to reconstruct the peatland. Testate amoebae
have been used to define the past nature of the peatland for
reconstruction purposes (Charman 1997; Charman &
Gilbert 1997).

Figure 52. Bullinularia indica. Photo by Edward Mitchell.

Jauhianinen (2002) demonstrated in an ombrotrophic

bog that the testacean shells were present throughout the
vertical profile, whereas in the minerotrophic fen they were

Chapter 2-5: Peatland Rhizopods

61

numerous only at the surface. As in other studies, moisture

conditions were important, but peat composition and
minerals also played important roles.
Following
restoration, species that indicated dry conditions
disappeared, whereas the moisture gradient seemed to
result in less defined community diffrences. In fact, the
minerals seemed to have a greater effect.

Figure 54. Opening of test of Euglypha compressa. Photo

by Edward Mitchell.

Figure 53. Nebela tincta test with living amoeba. Photo by

Edward Mitchell.

Lamentowicz et al. (2008) demonstrated that the

testate amoebae record in a Baltic coast peatland in
Northern Poland correlated well with the stable isotope
data in the same core. The large number of testate
protozoans known from peatlands (Table 1), their relatively
cosmopolitan distribution, and the understanding we have
of the water table requirements for many of these species
provide us with a useful tool for understanding the past
history of many peatlands.

62

Chapter 2-5: Peatland Rhizopods

Table 1. Species of testate amoebae known from peatlands. *Indictes species closely associated with Sphagnum.
Amphitrema flavum* - zoochlorellae
Amphitrema stenostoma * - zoochlorellae
Amphitrema wrightianum* - zoochlorellae
Arcella discoides*
Arcella gibbosa*
Arcella hemisphaerica*
Arcella mitrata
Arcella polypora
Arcella vulgaris*
Assulina muscorum*
Assulina seminulum*
Bullinularia indica*
Campascus minutus
Centropyxis aculeata group*
Centropyxis arcelloides*
Centropyxis cassis*
Corythion dubium*
Corythion pulchellum
Cryptodifflugia compressa
Cryptodifflugia eboracensis
Cryptodifflugia ovalis
Cryptodifflugia oviformis
Cryptodifflugia penardi
Cryptodiflugia pulex
Difflugia amphoralis
Difflugia bacilliarum*
Difflugia bacillifera*
Difflugia constricta
Difflugia curvicaulis
Difflugia globularis
Difflugia globulus
Difflugia oblonga*
Difflugia pyriformis
Difflugia rubescens*
Difflugia tuberculata*
Difflugia urceolata*
Euglypha ananthophora*
Euglypha brachiata
Euglypha ciliata*
Euglypha cristata
Euglypha filifera
Euglypha rotunda*
Euglypha scutigera
Euglypha strigosa*
Euglypha tuberculata*
Heleopera lata
Heleopera petricola*

Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Bovee 1979
Hingley 1993
Hingley 1993
Bovee 1979
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993
Hingley 1993

Heleopera rosea*
Hingley 1993
Heleopera sphagni* - zoochlorellae
Hingley 1993
Heleopera sylvatica*
Hingley 1993
Hyalosphenia cuneata
Hingley 1993
Hyalosphenia elegans* - zoochlorellae
Hingley 1993
Hyalosphenia minuta
Hingley 1993
Hyalosphenia ovalis
Hingley 1993
Hyalosphenia papilio* - zoochlorellae
Hingley 1993
Lecythium hyalinum
Hingley 1993
Lecythium mutabile
Hingley 1993
Lesquereusia epistomium
Hingley 1993
Lesquereusia inaequalis
Hingley 1993
Lesquereusia modesta*
Hingley 1993
Lesquereusia spiralis*
Hingley 1993
Nebela barbata*
Hingley 1993
Nebela bigibbosa*
Hingley 1993
Nebela carinata*
Hingley 1993
Nebela collaris*
Hingley 1993
Nebela dentistoma*
Hingley 1993
Nebela flabellum*
Hingley 1993
Nebela galeata*
Hingley 1993
Nebela griseola*
Hingley 1993
Nebela lageniformis*
Hingley 1993
Nebela marginata*
Hingley 1993
Nebela militaris*
Hingley 1993
Nebela minor*
Hingley 1993
Nebela parvula*
Hingley 1993
Nebela penardiana*
Hingley 1993
Nebela tenella
Mazei & Tsyganov 2007/08
Nebela tincta*
Gilbert et al. 2003
Nebela tubulosa*
Hingley 1993
Nebela vitraea*
Hingley 1993
Phryganella acropodia
Hingley 1993
Placocista jurassica
Hingley 1993
Placocista spinosa*
Hingley 1993
Portigulasia rhumbleri
Hingley 1993
Pseudochlamys patella
Hingley 1993
Quadrulella symmetrica*
Hingley 1993
Pseudodifflugia compressa
Hingley 1993
Pyxidicula cymbalum
Hingley 1993
Sphenoderia dentata
Hingley 1993
Sphenoderia fissirostris
Hingley 1993
Sphenoderia lenta*
Hingley 1993
Sphenoderia macrolepis
Hingley 1993
Trigonopyxis arcula*
Hingley 1993
Trinema enchelys*
Hingley 1993

Chapter 2-5: Peatland Rhizopods

Summary
Peatlands support an abundant bryophyte fauna,
with Amphitrema, Assulina, Corythion, Difflugia,
Euglypha, Heleopera, Hyalosphenia, and Nebela
typically being the most common genera. Sphagnum
sports more species than those found among other
mosses or tracheophytes. These taxa are widespread
and thus are very relaible indicators of moisture
conditions in the peatlands and are less affected by
water chemistry than are the tracheophytes.
Diversity is lowest in the driest peatland habitats,
but the number of individuals is highest. Abundance
increases with depth if oxygen is not limiting. Dry
habitat species are more tolerant of changes in water
depth than are wet habitat species. Rich fen amoeba
species differ from those of acid bogs, but Euglyphidae
are prominent in all these habitats. Paraquadrula
irregularis and Centropyxis discoides are restricted to
fens, with Arcella discoides indicative of rich fens.
Detritus forms a major portion of the protozoan diet in
the peatlands.
Vertical zonation presents the symbiotic taxa in the
light zone at the top of the moss, with those requiring
more moisture occurring at the greatest depths. Shell
size, pH, mosture, light, nutrients, and available food all
contribute to the distribution. Horizontal variation
results from differences in bryophyte species and
microtopography, resulting in differences in distance
from water table and in pH. Seasonal differences
reflect some of these same changes in moisture and
food availability and are effective in separating niches
of closely related species.
Amoebae form more constant associations in
peatlands than do the plants. And testate species, with
few exceptions, are well preserved even after death.
Therefore, they can serve as appropriate markers of past
climates as well as indicators of predisturbance
conditions, although tests of some species, especially
Euglyphidae, decompose more easily than others and
can skew the results. The best indicators are those with
narrow tolerance ranges, especially for moisture.

Acknowledgments
Edward Mitchell was particularly helpful in providing
me with needed pictures. Most of the others came from
photos by Yuuji Tsukii who gave me permission to use
anything of his on the Protist Information Server website.

Literature Cited
Bartos, E. 1949. Additions to knowledge of moss-dwelling fauna
of Switzerland. Hydrobiologia 2: 285-295.
Beyens, L. and Chardez, D. 1984. Testate amoebae (Rhizopoda,
Testaceae) from Southwest Ireland. Arch. Protistenkd. 128:
109-126.
Beyens, L. and Chardez, D. 1987. Evidence from testate
amoebae for changes in some local hydrological conditions
between c. 5000 BP and c. 3800 BP on Edgeoya (Svalbard).
Polar Res. 5: 165-169.
Beyens, L. and Meisterfeld, R. 2001. Protozoa: Testate
amoebae. In: Smol, J. P., Birks, H. J. B., and Last, W. M.
Tracking Environmental Change Using Lake Sediments.
Vol. 3. Terrestrial, Algal, and Siliceous Indicators. Kluwer
Academic Publishers, pp. 121-154.

63

Beyens, L., Chardez, D., Landtsheer, R. De, Bock, P. De, and

Jacques, E. 1986. Testate amoebae populations from moss
and lichen habitats in the Arctic. Polar Biol. 5: 165-173.
Bobrov, A. A., Charman, D. J., and Warner, B. G. 1999. Ecology
of testate amoebae (Protozoa: Rhizopoda) on peatlands in
western Russia with special attention to niche separation in
closely related taxa. Protistology 150: 125-136.
Bobrov, A. A., Charman, D. J., and Warner, B. G. 2002. Ecology
of testate amoebae from oligotrophic peatlands: Specific
features of polytypic and polymorphic species. Biol. Bull.
Rus. Acad. Sci. 29: 605-617.
Bonnet, L. 1958. Les thcamoebiens des Bouillouses. Bull. Soc.
Hist. Nat. Toulouse 93: 529-543.
Booth, R. K. 2001. Ecology of testate amoebae (protozoa) in two
Lake Superior coastal wetlands:
Implications for
paleoecology and environmental monitoring. Wetlands 21:
564-576.
Booth, R. K. 2002. Testate amoebae as paleoindicators of
surface-moisture changes on Michigan peatlands: modern
ecology and hydrological calibration. J. Paleolimnol. 28:
329-348.
Booth, R. and Jackson, S. 2001. A high-resolution record of late
Holocene surface-moisture changes from a Michigan raised
bog. Abstracts of the 86th meeting of the Ecological Society
of America, 6 August 2001.
Booth, R. K. and Zygmunt, J. R. 2005. Biogeography and
comparative ecology of testate amoebae inhabiting
Sphagnum-dominated peatlands in the Great Lakes and
Rocky Mountain regions of North America. Diversity
Distrib. 11: 577-590.
Booth, R. K., Jackson, S. T., and Gray, C. E. D. 2004.
Paleoecology and high-resolution paleohydrology of a kettle
peatland in upper Michigan. Quart. Res. 61: 1-13.
Bousquet, E. 1950. La faune sphagnicole (Thcamoebiens) de la
rgion d'Ax-les-Thermes (Arige). Annales du Laboratoire
d'Ax-les-Thermes 8: 39-46.
Bovee, E. C. 1979. Protozoa from acid-bog mosses and forest
mosses of the Lake Itasca region (Minnesota, U.S.A.). Univ.
Kans. Sci. Bull. 51: 615-629.
Buttler, A., Warner, B. G., Matthey, Y., and Grosvernier, P.
1996a. Testate amoebae (Protozoa: Rhizopoda) and
restoration of cut-over bogs in the Jura, Switzerland. New
Phytol. 134: 371-382.
Buttler, A., Warner, B. G., Grosvernier, P., and , and Matthey, Y.
1996b. Vertical patterns of testate amoebae (Protozoa:
Rhizopoda) and peat-forming vegetation on cutover bogs in
the Jura, Switzerland. New Phytol. 134: 371-382.
Chacharonis, P. 1956. Observations on the ecology of protozoa
associated with Sphagnum. J. Protozool. 3 (suppl.), Abstr.
58.
Charman, D. J. 1997. Modelling hydrological relationships of
testate amoebae (Protozoa: Rhizopoda) on New Zealand
peatlands. J. Royal Soc. N. Z. 27: 465-483.
Charman, D. J. and Warner, B. G. 1992. Relationship between
testate
amoebae
(Protozoa:
Rhizopoda)
and
microenvironmental parameters on a forested peatland in
northeastern Ontario. Can. J. Zool. 70: 2474-2482.
Charman, D. J. and Warner, B. G. 1997. The ecology of testate
amoebae (Protozoa: Rhizopoda) in oceanic peatlands in
Newfoundland, Canada:
Modelling hydrological
relationships for palaeoenvironmental reconstruction.
Ecoscience 4: 555-562.
Charman, D. J., Hendon, D., and Packman, S. 1999. Multiproxy
surface wetness records from replicate cores on an

64

Chapter 2-5: Peatland Rhizopods

ombrotrophic mire: Implications for Holocene palaeoclimate

records. J. Quatern. Sci. 14: 451-463.
Charman, D. J., Hendon, D., and Woodland, W. 2000. The
identification of peatland testate amoebae. Quaternary
Research Association Technical Guide London 9: 147.
Conrad, W. 1943. Sur quelques organismes sphagnicoles.
Bulletin du Museum d'Histoire Naturelle de Belgique 19: 14.
Corbet, S. A. 1973. An illustrated introduction to the testate
rhizopods in Sphagnum, with special reference to the area
around Malham Tarn, Yorkshire. Field Stud. 3: 801-838.
Coteaux, M.-M. 1969. Thcamoebiens muscicoles de Gaume et
de Moyenne-Belgique. Rev. Ecol. Biol. Sol. 6: 413-428.
Coteaux, M.-M. 1992. Decomposition of cells and empty shells
of testate amoebae (Rhizopoda, Testacea) in an organic acid
soil sterilized by propylene oxide fumigation, autoclaving,
and -ray irradiation. Biol. Fert. Soils 12: 290-294.
Douglas, M. S. V. and Smol, J. P. 1988. Siliceous protozoan and
chrysophycean microfossils from the recent sediments of
Sphagnum dominated Lake Colden, N.Y., USA.
In:
Sladecek, V. (ed.).
Proceedings of the International
Association of Theoretical and Applied Limnology,
Congress in New Zealand (Hamilton (New Zealand)), 8 Feb
1987 Ann. Zool. Soc. Zool.-Bot. Fenn. "Vanamo" 23(2):
855-859.
Douglas, M. S. V. and Smol, J. P. 2001. Siliceous protozoan
plates and scales. Chapt. 13. In: Smol, J. P., Birks, H. J.
B., Last. (eds.).
Tracking Environmental Change Using
Lake Sediments. Vol. 3: Terrestrial, Algal, and Siliceous
Indicators. Kluwer Academic Publishers, Dordrecht, The
Netherlands.
Foissner, W. 1999. A Holocene record of climate, vegetation
change and peat bog development, east Otago, South Island,
New Zealand. J. Quatern. Sci. 14: 239-254.
Geel, E. van. 1976. A paleontological study of Holocene peat
bog sections based on the analysis of pollen, spores and
micro- and microscopic remains of fungi, algae, cormophytes
and animals. Hugo de Vries Laboratorium, Universiteit van
Amsterdam (Amsterdam), pp. 1-75.
Gerson, U. 1982. Bryophytes and invertebrates. In: Smith, A. J.
E. (ed.). Bryophyte Ecology. Chapman & Hall, New York.
Pp. 291-332.
Gilbert, D. and Mitchell, E. A. D. 2006. Microbial diversity in
Sphagnum peatlands. In: Martini, I. P., Martinez Cortizas,
A., Chesworth, W. (Eds.). Peatlands: Evolution and
Records of Environmental and Climate Changes. Elsevier B.
V. Chapt. 13, pp. 287-317.
Gilbert, D., Amblard, C., Bourdier, G., and Francez, A.-J. 1998a.
The microbial loop at the surface of a peatland: Structure,
function, and impact of nutrient input. Microb. Ecol. 35: 8393.
Gilbert, D., Amblard, C., Bourdier, G., and Francez, A.-J. 1998b.
Short-term effect of nitrogen enrichment on the microbial
communities of a peatland. Hydrobiologia 374: 111-119.
Gilbert, D., Mitchell. E. A. D., Amblard, C., Bourdier, G., and
Francez, A.-J. 2003. Population dynamics and food
preferences of the testate amoeba Nebela tincta majorbohemica-collaris complex (protozoa) in a Sphagnum
peatland. Acta Protozool. 42: 99-104.
Grospietsch, T. 1976. Rhizopodenanalytische Untersuchungen.
In Zwei Mooren im oberbayerischen Jungmornengebiet
sdlich des Starnberger Sees (Weidfilz und Schgenfilz).
Telma (Hanover) 6: 25-34.
Harnish, O. 1924. Studien zur Okologie der Moorfauna. Biol.
Zentralbl. 44: 110-127.

Harnish, O. 1925. Studien zur kologie und Tiergeographie der

Moore. Zoologisch Jahrbuch (Abteilung Systematik) 51: 1166.
Harnish, O. 1927. (1929??). Einige Daten zur rezenten und
fossilen testaceen Rhizopodenfauna der Sphagnen. Arch.
Hydrobiol. 18: 345-360.
Harnish, O. 1948. Rhizopodenanalyse der Moore. Biol.
Zentralbl. 67: 551-562.
Harnish, O. 1950. Daten zur Gestaltung der oekologischen
Valenz der sphagnicolen Rhizopoden im Verlauf des
Postglazials. Deutsche Zoologische Zeitschrift (Hannover)
1: 222-233.
Harnish, O. 1951. Rhizopodenanalytische Studien an einigen
Vogesenmooren. Arch. Hydrobiol. 45: 332-345.
Heal, O. 1961. The distribution of testate amoeba (Rhizopoda,
Testacea) in some fens and bogs in northern England. J.
Linn. Soc. Zool. 44: 369-382.
Heal, O. W. 1964. Observations on the seasonal and spatial
distribution of Testacea (Protozoa: Rhizopoda) in Sphagnum.
J. Anim. Ecol. 33: 395-412.
Heinis, F.
1945.
Beitrge zur Mikrobiozonose der
Sphagnumpolster auf God del Fourn im Nationalpark.
Ergebnisse Wissenschaften Unters. Schwiez. Nationalparkes
N.S. 1: 527-547.
Hendon, D. and Charman, D. J. 1997. The preparation of testate
amoebae (Protozoa:
Rhizopoda) samples from peat.
Holocene 7: 199-205.
Hingley, M. 1993. Microscopic Life in Sphagnum. Illustrated by
Hayward, P. and Herrett, D. Naturalists' Handbook 20. [iiv]. Richmond Publishing Co. Ltd., Slough, England, 64 pp..
58 fig. 8 pl. (unpaginated).
Hoogenraad, H. R. 1934. Studien ber die sphagnicolen
Rhizopoden der niederlndischen Fauna. D. Sci. thesis,
University of Utrecht, Lippert and Co. (Naumburg - Saale),
pp. I-IV + 1-100.
Hoogenraad, H. R. 1935. Studien ber die sphagnicolen
Rhizopoden der niederlndischen Fauna. Arch. Protistenk.
84: 1-100.
Hoogenraad, H. R. and Groot, A. A. de. 1940. Moosbewohnende
thekambe Rhizopoden von Java und Sumatra. Treubia
(Buitenzorg) 17: 209-259.
Hoogenraad, H. R. and Groot, A. A. de. 1946. Thekamoebe
sphagnumrhizopoden van Buitenzorg (Java). Biologische
Jaarboek, Dodonea (Gent) 13: 112-126.
Hoppman, J. 1954. Die Thekamoeben der Tormoose des venner
Moores. Nature und Heimat 14: 1-8.
Hughes, P. D. M., Blundell, A., Charman, D. J., Bartlett, S.,
Daniell, J. R. G., Wojatschke, A., and Chambers, F. M.
2006. An 8500 cal. year multi-proxy climate record from a
bog in eastern Newfoundland: Contributions of meltwater
discharge and solar forcing. Quart. Sci. Rev. 25: 1208-1227.
Jackzo, I. 1941. Nhany dunantuli atmeneti tozegmohalap s
Sphagnum elofordulas hazas Rhizopodairol.
Allatani
Kzlemnyek 38: 18-34.
Jauhiainen, S. 2002. Testacean amoebae in different types of
mire following drainage and subsequent restoration. Eur. J.
Protistol. 38: 59-72.
Jung, W. 1936. Thekamoben eines Eggegebirgsmoores und
zweier Moore im Hohen Venn. Ann. Protistol. 5: 83-123.
Kishaba, K. and Mitchell, E. A. D. 2005. Changes in testate
amoebae (protists) communities in a small raised bog. A 40year study. Acta Protozool. 44: 1-12.
Laggoun-Dfarge, F., Mitchell, E., Gilbert, D., Disnar, J. R.,
Comont, L., Warner, B. G., and Buttler, A. 2008. Cut-over

Chapter 2-5: Peatland Rhizopods

peatland regeneration assessment using organic matter and

microbial indicators (bacteria and testate amoebae). J. Appl.
Ecol. 42: 716-727.
Lamentowicz, M. and Mitchell, E. A. D. 2005. The ecology of
testate amoebae (protists) in Sphagnum in north-western
Poland in relation to peatland ecology. Microbial Ecol. 50:
48-63.
Lamentowicz, L., Gbka, M., and Lamentowicz, M. 2007.
Species composition of testate amoebae (protists) and
environmental parameters in a Sphagnum peatland. Polish J.
Ecol. 55: 49-59.
Lamentowicz, M., Cedro, A., Gaka, M., Goslar, T., MiotkSzpiganowicz, G., Mitchell, E. A. D., and Pawlyta, J. 2008.
Last millennium palaeoenvironmental changes from a Baltic
bog (Poland) inferred from stable isotopes, pollen, plant
macrofossils and testate amoebae.
Palaeogeogr.
Palaeoclimatol. Palaeoecol. 265: 93-106.
Lamentowicz, M., Tobolski, K., and Mitchell, E. A. D. 2007.
Palaeoecological evidence for anthropogenic acidification of
a kettle-hole peatland in northern Poland. Holocene 17:
1185-1196.
Laminger, H. 1975. Die Sukzession der Testaceen-Assoziationen
(Protozoa, Rhizopoda) im rezenten und subfossilen
Sphagnum des Obersees bei Lunz (Niedersterreich). [The
succession of Testacea-associations (Protozoa, Rhizopoda) in
recent and subfossil Sphagnum from the Obersee near Lunz
(Austria).]. Hydrobiologia 46: 465-487.
Langdon, P. G., Barber, K. E., and Hughes, P. D. M. 2003. A
7500-year peat-based palaeoclimatic reconstruction and
evidence for an 1100-year cyclicity in bog surface wetness
from Temple Hill Moss, Pentland Hills, Southeast Scotland.
Quart. Sci. Rev. 22: 259-274.
Leidy, J. 1879. On rhizopods occurring in Sphagnum. Proc.
Acad. Nat. Sci. Philadelphia 3rd ser. 31: 162-163.
Martin, W.
1963.
Zooznologische Analyse einer
Waldsphagnetum aus dem Erzgebirge. Wissenschaftliche
Beitraege, Inst. Mhlhausen (Thr.) 1: 77-95.
Mattheeussen, R., Ledeganck, P., Vincke, S., Vijver, B. van de,
Nijs, I. and Beyens, L. 2005. Habitat selection of aquatic
testate amoebae communities on Qeqertarsuaq (Disco
Island), West Greenland. Acta Protozool. 44: 253-263.
Mauquoy, D. and Barber, K. 2002. Testing the sensitivity of the
palaeoclimatic signal from ombrotrophic peat bogs in
northern England and the Scottish Borders. Rev. Palaeobot.
Palynol. 119: 219-240.
Mazei, Y. A. and Bubnova, O. A. 2007. Species composition and
structure of testate amoebae community in a Sphagnum bog
at the initial stage of its formation. Biol. Bull. 37: 619-628.
[Orig.: Izvestiya Akademii Nauk, Seriya Biologicheskaya,
2007, No. 6, pp. 738747.].
Mazei, Y. A. and Tsyganov, A. N.
2007/08.
Species
composition, spatial distribution and seasonal dynamics of
testate amoebae community in a Sphagnum bog (Middle
Volga region, Russia). Protistology 5: 156-206.
Mazei, Y. A., Tsyganov, A. N., and and Bubnova, O. A. 2007.
Structure of a community of testate amoebae in a Sphagnum
dominated bog in upper Sura Flow (Middle Volga Territory).
Biol. Bull. 34: 382-394.
McGlone, M. S. and Wilmshurst, J. M. 1999a. A Holocene
record of climate, vegetation change and peat bog
development, east Otago, South Island, New Zealand. J.
Quatern. Sci. 14: 39-254.
McGlone, M. S. and Wilmshurst, J. M. 1999b. Multiproxy
surface wetness records from replicate cores on an
ombrotrophic mire: Implications for Holocene palaeoclimate
records. J. Quatern. Sci. 14: 451-463.

65

Meisterfeld, R. 1977. Die horizontale und vertikale Verteilung

der Testaceen (Rhizopoda, Testacea) in Sphagnum. Arch.
Hydrobiol. 79: 319-356.
Meisterfeld, R. 1978. Die Struktur von Testaceenznosen
(Rhizopoda, Testacea) in Sphagnum unter besonderer
Bercksichtigung ihrer Diversitt. Verh. Gesell. kol. 7:
441-450.
Meisterfeld, R.
1979a.
Zur Systematik der Testaceen
(Rhizopoda, Testacea) in Sphagnum. Eine REMUntersuchung. Arch. Protistenk. 121: 246-269.
Meisterfeld, R. 1979b. Clusteranalytische Differenzierung der
Testaceenznosen (Rhizopoda, Testacea) in Sphagnum.
Arch. Protistenk. 121: 270-307.
Meisterfeld, R. and Heisterbaum, M. 1986. The decay of empty
tests of testate amoebae (Rhizopoda, Protozoa). Symp. Biol.
Hung. 33: 285-390.
Mieczan, T. 2006. Species diversity of protozoa (Rhizopoda,
Ciliata) on mosses of Sphagnum genus in restoration areas of
the Poleski National Park. Acta Agrophys. 7: 453-459.
Mieczan, T. 2007. Epiphytic protozoa (testate amoebae and
ciliates) associated with Sphagnum in peatbogs:
Relationship to chemical parameters. Polish J. Ecol. 55: 7990.
Mitchell, E. A. D. 2004. Response of testate amoebae (protozoa)
to N and P fertilization in an Arctic wet sedge tundra. Arct.
Antarct. Alp. Res. 36: 77-82.
Mitchell, E. A. D. and Gilbert, D. 2004. Vertical microdistribution and response to nitrogen deposition of testate
amoebae in Sphagnum. J. Eukaryot. Microbiol. 51: 480-490.
Mitchell, E. A. D. and Meisterfeld, R. 2005. Taxonomic
confusion blurs the debate on cosmopolitanism versus local
endemism of free-living protists. Protist 156: 263-267
Mitchell, E. A. D., Borcard, D., Buttler, A. J., Grosvernier, Ph.,
Gilbert, D., and Gobat, J.-M. 2000a. Horizontal distribution
patterns of testate amoebae (protozoa) in a Sphagnum
magellanicum carpet. Microbiol. Ecol. 39: 290-300.
Mitchell, E. A. D., Buttler, A. J., Warner, B. G., and Gobat, J. M.
1999b. Ecology of testate amoebae (Protozoa: Rhizopoda)
in Sphagnum peatlands in the Jura mountains, Switzerland
and France. Ecoscience 6: 565-576.
Mitchell, E. A. D., Buttler, A., Grosvernier, P., Rydin, H.,
Albinsson, C., Greenup, A. L., Heijmans, M. M. P. D.,
Hoosbeek, M. R., and Saarinen, T. 2000b. Relationships
among testate amoebae (protozoa), vegetation and water
chemistry in five Sphagnum-dominated peatlands in Europe.
New Phytol. 145: 95-106.
Mitchell, E. A. D., Charman, D. J., and Warner, B. G. 2008.
Testate amoebae analysis in ecological and paleoecological
studies of wetlands: past, present and future. Biodivers.
Conserv. 17: 2115-2137.
Mitchell, E. A. D., Gilbert, D., Buttler, A., Amblard, C.,
Grosvernier, P., and Gobat, J.-M. 2003. Structure of
microbial communities in Sphagnum peatlands and effect of
atmospheric carbon dioxide enrichment. Microbial Ecol.
46: 187-199.
Mitchell, E. A. D., Knaap, W. O. van der, Leeuwen, J. F. N. van,
Buttler, A., Warner, B. G., and Gobat, J.-M. 2001. The
palaeoecological history of the Praz-Rodet bog (Swiss Jura)
based on pollen, plant macrofossils and testate amoebae
(Protozoa). Holocene 11: 65-80.
Mitchell, E. A. D., Payne, R. J., and Lamentowicz, M. 2007.
Potential implications of differential preservation of testate
amoeba shells for paleoenvironmental reconstruction in
peatlands. J. Paleolimnol. 40: 603-618.

66

Chapter 2-5: Peatland Rhizopods

Nguyen-Viet, H., Gilbert, D., Bernard, N., Mitchell, E. A. D., and

Badot, P.-M. 2004. Relationship between atmospheric
pollution characterized by NO2 concentrations and testate
amoebae density and diversity. Acta Protozool. 43: 233-239.
Nguyen-Viet, H., Gilbert, D., Mitchell, E. A. D., Badot, P.-M.,
and Bernard, N. 2007. Effects of experimental lead
pollution on the microbial communities associated with
Sphagnum fallax (Bryophyta). Mycrobial Ecol. 54: 232-241.
Nguyen-Viet, H., Bernard, N., Mitchell, E. A. D., Badot, P.-M.,
and Gilbert, D. 2008. Effect of lead pollution on testate
amoebae communities living in Sphagnum fallax: An
experimental study. Ecotoxicol. Environ. Safety 69: 130138.
Opravilov, V. and Hjek, M. 2006. The variation of testacean
assemblages (Rhizopoda) along the complete base-richness
gradient in fens: A case study from the Western Carpathians.
Acta Protozool. 45: 191-204.
Opravilov, V. and Zahrdkov, S. 2003. Some information of
the testate amoebae of Iceland. Limnologica 33: 131-137.
Oye, P. van. 1941. Die Rhizopoden des Sphagnetums bei
Krisuvik auf Island. Biologisch Jaarboek Dodonaea (Gent)
8: 284-305.
Oye, P. van. 1951. Die Rhizopoden des Sphagnetums bei
Krisuvik auf Island. Biologische Jaarboek, pp. 284-327.
Paulson, B. 1952-1953. Some rhizopod associations in a
Swedish mire. Oikos 4: 151-165.
Payne, R. 2007. Laboratory experiments on testate amoebae
preservation in peats: implications for palaeoecology and
future studies. Protozoology 46: 325-332.
Payne, R. J. and Mitchell, E. A. D. 2007. Ecology of testate
amoebae from mires in the Central Rhodope Mountains,
Greece and development of a transfer function for
palaeohydrological reconstruction. Protist 158: 159-171.
Payne, R. J., Charman, D. J., Matthews, S., and Eastwood, W. J.
2008. Testate amoebae as palaeohydrological proxies in
Srmene Aabai Yaylasi Peatland (Northeast Turkey).
Wetlands 28: 311-323.
Payne, R. J., Kishaba, K., Blackford, J. J., and Mitchell, E. A. D.
2006. Ecology of testate amoebae (Protista) in south-central
Alaska peatlands: Building transfer-function models for
palaeoenvironmental studies. Holocene 16: 403-414.
Robson, T., Ballar, C., Sala, O., Scopel, A., and Caldwell, M.
2001. Biodiversity of microfauna and fungal communities in
a Sphagnum bog under two levels of solar UV-B. Abstracts
of the 86th meeting of the Ecological Society of America,
Madison, WI, 3-8 August 2001.
Rose, F. 1953. A survey of the ecology of the British lowland
bogs. Proc. Linn. Soc. London 164: 186-211.
Ruitenburg, G. J. and Davids, C. 1977. Thecamoeba in
succession series of some peat-bog vegetations. Hydrobiol.
Bull. (Amsterdam) 2: 22-24.
Schnitchen, C., Magyari, E., Tthmrsz, B, Grigorszky, I., and
Braun, M. 2003. Micropaleontological observations on a
Sphagnum bog in East Carpathian region testate amoebae
(Rhizopoda: Testacea) and their potential use for
reconstruction of micro- and macroclimatic changes.
Hydrobiologia 506: 45-49.
Schnborn, W. 1962. Zur kologie der sphagnikolen, bryokolen
und terrikolen Testaceen. Limnologica 1: 231-254.
Schnborn, W. 1963. Die Stratigraphie lebender Testaceen im
Sphagnetum der Hochmoore. Limnologica 1: 315-321.
Schnborn, W. 1965. Untersuchungen ber die ZoochlorellenSymbiose der Hochmoor-Testaceen. Limnologica 3: 173176.

Schoning, K., Charman, D. J., and Wastegoard, S.

2005.
Reconstructed water tables from two ombrotrophic mires in
eastern central Sweden compared with instrumental
meteorological data. Holocene 15: 111-118.
Seis, J. 1971. Rhizopodenanalytische untersuchungen an den
mooren des pleistozaenen Salzachvorlandgletschers,
Vorbericht, Leopoldskroner Moor. Berichte aus dem Haus
der Natur in Salzburg, Abteilung B, GeologischMineralogische Sammlungen 2: 10-14.
Smith, H. G. and Wilkinson, D. M. 2007. Not all free-living
microorganisms have cosmopolitan distributions the case
of Nebela (Apodera) vas Certes (Protozoa: Amoebozoa:
Arcellinida). J. Biogeogr. 34: 1822 1831.
Strder-Kypke, M. C. and Schnborn, W. 1999. Periphyton and
sphagnicolous protists of dystrophic bog lakes (Brandenburg,
Germany): II. Characteristic species and trophy of the lakes.
Swindles, G. T. and Roe, H. M. 2007. Examining the dissolution
characteristics of testate amoebae (Protozoa: Rhizopoda) in
low pH conditions: Implications for peatland palaeoclimate
studies. Palaeogeogr. Palaeoclimatol. Palaeoecol. 252: 486496.
Thomas, R. 1959. Les Thcamoebiens muscicoles et terricoles:
Notions d'cologie gnrale et comparative. Procs-Verbaux
de la Socit Linnenne de Bordeaux 98: 27-53.
Tolonen, K. 1966. Stratigraphic and rhizopod analyses on an old
raised bog, Varrassuo, in Hollola, South Finland. Ann. Bot.
Fenn. 3: 147-166.
Tolonen, K. 1994. Ecology of testaceans (Protozoa, Rhizopoda)
in Mires in Southern Finland.2. Multivariate-analysis. Arch.
Protistenk. 144: 97-112.
Tolonen, K., Huttunen, P., and Jungner, H. 1985. Regeneration
of two coastal raised bogs in eastern North America:
Stratigraphy, radiocarbon dates and rhizopod analysis from
sea cliffs. Ann. Acad. Sci. Fenn., Ser. A, III. GeologicaGeographica 139: 5-51.
Tolonen, K., Warner, B. G., and Vasander, H. 1992. Ecology of
testaceans (Protozoa: Rhizopoda) in mires in southern
Finland: I. Autecology. Arch. Protistenk. 142: 119-138.
Tolonen, K., Warner, B. G., and Vasander, H. 1994. Ecology of
testaceans in mires in southern Finland: II. Multivariate
analysis. Arch. Protistenk. 144: 97-112.
Varga, L. 1956. Adatok a hazai Sphagnum lapok vizi
mikrofaunajanaki ismerethez. Allatani Kslemnyek 16:
149-158.
Vincke, S., Gremmen, N., Beyens, L., and Vijver, B. Van de.
2004. The moss dwelling testacean fauna of le de la
Possession. Polar Biol. 27: 753-766.
Vucetich, M. C. 1975. Tecamebianos musccolas y esfagncolas
de islas Malvinas (Argentina). Neotropica 21: 11-16.
Warner, B. G. 1987. Abundance and diversity of testate amoebae
(Rhizopoda, Testacea) in Sphagnum peatlands in
southwestern Ontario, Canada. Arch. Protistenk. 133: 173189.
Warner, B. G. 1991. Fossil testate amoebae (protozoa) and
hydrological history of an ombrotrophic bog in northwestern
Ontario, Canada. In: Spigarelli, S.A. (ed.). Proceedings of
an International Symposium on Peat/Peatland Characteristics
and Uses. Minnesota, Bemidji State University, pp. 5-14.
Warner, B. G. and Charman, D. J. 1994. Holocene changes on a
peatland in northwestern Ontario interpreted from testate
amoebae (protozoa) analysis. Boreas 23: 270-279.
Warner,
Warner, B. G., Asada, T., and Quinn, N. P. 2007. Seasonal
influences on the ecology of testate amoebae (protozoa) in a

Chapter 2-5: Peatland Rhizopods

small Sphagnum peatland in Southern Ontario, Canada.

Microbiol. Ecol. 54: 91-100.
Wilmshurst, Janet M.
1998.
And in the bog there
lived...Protozoan bog dwellers can tell us a lot about past
climates. New Zealand Science Monthly Online. Accessed
on
28
October
2008
at
<http://nzsm.webcentre.co.nz/article1767.htm>.
Woodland, W. A. 1998. Quantitative estimates of water tables
and soil-moisture in Holocene peatlands from testate amebas.
Holocene 8: 261-273.
Woodland, W. A., Charman, D. J., and Sims, P. C. 1998.
Quantitative estimates of water tables and soil moisture in
Holocene peatlands from testate amoebae. Holocene 8: 261273.
Zygmunt, J., Booth, R., and Jackson, S. 2003. Ecology of testate
amoebae in Rocky Mountain peatlands and their application
to paleohydrological reconstruction. Abstracts of the 88th
meeting of the Ecological Society of America, Savannah,
GA, 3-8 August 2003 Accessed on 3 July 2004 at
<http://abstracts.co.allenpress.com/pweb/esa2003/document/
?ID=26603>.

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